REVIEW
Acta Theriologica 45 (3): 289-307, 2000.
P L ISSN 0 0 0 1 - 7 0 5 1
Valid and invalid nomenclature of living and fossil deer,
Cervidae
Peter GRUBB
Grubb P. 2000. Valid and invalid nomenclature of living and fossil deer, Cervidae. Acta
Theriologica 45: 289-307.
Investigation of cervid nomenclature has revealed unavailable or preoccupied
names still in use; unnoticed or unevaluated homonymy; unused or unnoticed names,
including senior synonyms; unnoticed or misidentified types of genera; mis-cited
authorship; unjustified emendations of original spelling; and corrections of nomenclatural errors that have been neglected in subsequent literature. The following names
appearing in recent literature are affected: Pliocervinae Khomenko, Neocervinae,
Cervulinae (unavailable names); Capreolinae, Alceinae, Rangiferinae (attributable to
Brookes, 1828, not to authors who changed their rank or corrected original spelling;
take precedence over Odocoileinae when the taxa are combined, contrary to common
practice); Alcinae (emendation due to Blyth, not Jerdon, now superseded by Alceinae,
with priority over Rangiferinae - where relevant - here designated); Muntiacinae
(author is Knottnerus-Meyer, 1907, not Pocock, 1923; Elaphodinae here designated a
junior synonym); Megacerinae Viret (preoccupied by Megaloceridae Brookes, emended
to Megalocerotinae); Blastocerus Wagner (an available name of which Blastóceros
Fitzinger is an unjustified emendation, not a senior synonym of Ozotoceros\ lectotype
confirmed to be Ceruus paludosus Desmarest, 1822); Dorcelaphus (junior synonym of
Odocoileus, not a senior synonym of Blastocerus); Mazama gouazoubira,
Muntiacus
feai, Pudu pudu (unjustified emendations); Ceruus japonicus Otsuka, 1967 (preoccupied,
new name proposed); Cervus elaphus montanus Botezat, 1903 (nomen nudum and
preoccupied); and Pseudodarna (preoccupied by Metacervocerus).
35 Downhills Park Road, London N17 6PE, U.K.
Key words: Cervidae, deer, nomenclature
I n t r o d u c t i o n and m e t h o d s
Problems concerning the nomenclature of cervids are still current and the
following account, based on a review of the literature, deals with these issues. No
attempt has been made to address misspellings, such as Capriolinae, Alcenae,
Odocoilenae, Odocoilinae and Odocoilini.
H o l o m e t a c a r p a l or p l i o c e r v i n e d e e r
Pliocervinae Khomenko, 1914, is not an available name, having been coined to
include only Cervavitus, Cervocerus and Damacerus, all of Khomenko, 1913, and
[289]
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P. Grubb
therefore not based on a genus-group name (ICZN 1985: Article 29). After the
subsequent description of a genus Pliocervus by Hilzheimer (1922), Teilhard de
Chardin and Trassert (1937) re-diagnosed Pliocervinae Khomenko but did not
validate the name, as they included only Cervocerus and Cervavitus in the
subfamily. Viret (1961) omitted Pliocervus from his account of the Pliocervinae.
Czyżewska (1968) reduced Pliocervinae to a tribe, Pliocervini, to include Cervavitus,
but cited Pliocervus as incertae sedis within this tribe! Pliocervini Khomenko is still
being cited as if it was an available name (Vislobokova 1990).
Pliocervinae Symeonidis, 1974: 311, is an available name which can be taken to
be based on Pliocervus Hilzheimer, 1922 (McKenna and Bell 1997).
Cervavitus tarakliensis, Cervocerus novorossiae and Damacerus bessarabiae, all
of Khomenko, 1913: 108-109 and 133-134, are based on two-tined, three-tined and
palmate antlers, respectively. Zdansky (1925: 12) included Damacerus in the
synonymy of Cervocerus, hence acting as first reviser (ICZN 1985: Article 24), and
his synonymy was followed by Simpson (1945: 154). The relative rank of Cervavitus
and Cervocerus remained to be decided. Pidoplitschko and Flerov (1952: 124)
followed by Czyżewska (1968) included Cervoceros [sicl and Damaceros [sic] in the
synonymy of Cervavitus, so finalising the relative priority of Khomenko's names.
Azzaroli (1953: 9) recognised the work of Zdansky (1925) yet included Ceravitus
and Cervocerus in the synonymy of Damacerus, action which cannot be accepted in
view of the previously established priorities.
Procervus Alexejev, 1913: 1, with type by monotypy P. variabilis Alexejev, loc.
cit., is preoccupied by Procervus de Blainville, 1840 (a synonym of Rangifer
Hamilton Smith, 1827), and Procervus Hodgson, 1847 (a synonym of Cervus
(Rucervus) Hodgson, 1838). Azzaroli (1953) treated Procervus Alexejev as a
synonym of Damacerus (= Cervavitus). Viret (1961) still cited Procervus Alexejev
as a genus distinct from Cervocerus but therefore should have employed the name
Metadicrocerus.
Metadicrocerus Schlosser, 1924: 164 ('Metadircocerus' on p. 75), is a neglected
substitute for Procervus Alexejev, 1913, which was regarded as an unsuitable name.
Pliocervus Hilzheimer, 1922: 743, was proposed as a new genus with type Cervus
matheronis Gervais, 1852, by monotypy. Although regarded by Kretzoi (1941,
1968) as preoccupied, this is not the case.
Ctenocerus Kretzoi, 1941: 351, with type Cervus matheronis Gervais, 1852, was
substituted for Pliocervus Hilzheimer, which was said to be preoccupied by
Pliocervinae Khomenko, 1913, though an unavailable family-group name can not
threaten a genus-group name. As Pliocervinae Khomenko is unavailable, Pliocervus
should stand. In any case, Ctenocerus Kretzoi is preoccupied by Ctenocerus
Dahlbom, 1845 (Hymenoptera; Kretzoi 1968: 164) though remains a junior objective
synonym of Pliocervus Hiltzheimer.
Ctenocervus Kretzoi, 1968: 164, is a substitute for Ctenocerus Kretzoi, preoccupied, and therefore is another junior objective synonym of Pliocervus Hiltzheimer.
Nomenclature of living and fossil deer
291
Antlered t e l e m e t a c a r p a l deer
Telemetacarpi Brooke, 1878: 915, is unavailable as it is not based on a generic
name (ICZN 1985: Article 35). It is equivalent to the available name Capreolinae in
the sense of Pocock (1910: 971). Adjectival derivatives of telemetacarpi and of the
related plesiometacarpi have included telemetacarpalid and plesiometacarpalid
(Hershkovitz 1982: 9) or telemetacarpaline and plesiometacarpaline (Gustafson
1985: 89), but telemetacarpal and plesiometacarpal may suffice. Pocock (1910) used
the terms Telemetacarpalia and Plesiometacarpalia.
Capreolidae Brookes, 1828: 62, was coined as a subfamily of Cervidae, with type
genus Capreolus Gray, 1821, by definition, so when correctly formed is Capreolinae:
Gray (1852: x) is the author of this spelling. The name was first proposed at the same
time as Alcedae (Alceinae) and Rangiferinidae (Rangiferinae), but may be regarded
as the senior family-group name for antlered telemetacarpal deer through first-reviser action of Pocock (1910: 971). Simpson (1945: 267) referred to the 'group,
sometimes called Capreolinae for historic reasons but here called Odocoileinae
because Odocoileus is a better type than Capreolus ...'. He included Capreolini
Brookes, 1828, with a new rank as a tribe within the subfamily Odocoileinae
Pocock, 1923, but the reverse is correct: Odocoileini within Capreolinae. 'Capreolini
Simpson' is cited by Viret (1961), but Simpson authored the change in rank, not the
name. Cited as 'Capriolinae [sic] Pocock, 1910' as a synonym of Neocervinae by
Banfield (1961), but Pocock is not even the author of a correctly emended spelling,
let alone of the name itself.
Alcedae Brookes, 1828: 61, was coined as a subfamily of Cervidae with type
genus Alces Gray, 1821, by definition, with what would now be recognised as an
incorrectly formed suffix. Alceae (Gray 1852: 186) is a subtribal name but equivalent
to a subfamily, based on Alces. Brookes' name is acknowledged, so Alceae may be
regarded as an emendation of Alcedae, but the suffix is still incorrectly formed.
Alcadae (Gray 1872: 66; Gray 1873: 136) is a family name based on Alces yet with
incorrect generic stem. No reference to Brookes (1828) was made but it may be
regarded as a further emendation of Alcedae Brookes. It is a homonym of another
improperly formed name, Alcadae Anonymous, 1820, based on Alca Linnaeus, 1758
(the razorbill A. torda\ Aves; Kashin 1974). 'Alcinae Jerdon, 1874: 253' is quoted by
Palmer (1904: 726), Simpson (1945: 155) and Haltenorth (1963: 48) but this is not a
new name or even designation of a new rank, only an emendation, and it is due to
Blyth (1863: 145), not Jerdon. The suffix is correctly formed and the prefix - the
generic stem - has proved acceptable, for as a subfamily 'Alcinae' (or a tribe, Alcini)
has been widely used. But as Kashin (1974) has pointed out, it is a homonym of
family-group names based on Alca by justifiable emendation of Alcadae Anonymous,
1820, to Alcidae, Alcinae and Alcini. Both have the same stem Ale-. At Kashin's
instigation, the Commission has ruled that 'Alceinae' is now to be the approved
emendation of Alcedae Brookes (ICZN 1977b: Opinion 1081), with stem Alee-.
Contrary to the views of Eisenmann, Tortonese or Kraus (ICZN 1977b), a family-
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P. Grubb
-group name based on Alces (if not a family) has been in use for over 170 years, as
demonstrated above. 'Alcini Simpson' is cited by Viret (1961), but Simpson
authored the change in rank, not the name. If Capreolus is to be excluded from a
group otherwise containing Alces, Rangifer and genera of deer confined to the
Americas, called the Odocoileidae by Bubenik (1990), then the prior name should
be either Alceidae or Rangiferidae. The relative priority of these two names has not
been established. In addressing this problem, Alceinae is here designated as having
seniority in synonymy over Rangiferinae, the present author acting as first reviser.
Rangiferinidae Brookes, 1828: 61, was coined as a subfamily of Cervidae based
on Rangifer Hamilton Smith, 1827, and hence is now seen to have improperly
constructed prefix and suffix. Correctly the stem is Rangifer-, so the name is
properly Rangiferinae. Gray (1852: 188) emended Rangiferinidae to Rangerinae
and then (Gray 1872: 66) as a family to Rangiferidae. Rangiferinae is cited as 'New
Subf. Bubenik 1986c (= Rangiferidae Brookes, 1828)' by Bubenik (1990) but
subfamily rank is attributable to the original author of the name, correctly
amended spelling of the name to Gray (1872) and first correct formation of the
subfamily name to Pocock (1923). Viret (1961) cited 'Rangiferini Simpson' but
Simpson (1945) authored the change in ranking, not the name. Rangiferinae has
priority over Odocoileinae if the two taxa are combined.
Elaphalcedae Brookes, 1828: 62, is a subfamily name with incorrectly formed
suffix, based on Elaphalces Brookes, loc. cit., and with type species E. mexicanus (a
quotation of Ceruus mexicanus Gmelin, 1788, = Odocoileus virginianus
mexicanus).
Brookes termed the genus Elaphalces 'Gouazou Pougou', a name cited by Azara for
the marsh deer, Blastocerus dichotomous, and gave it the vernacular qualifying
epithet 'Mexican'. The presumed synonymy of the gouazoupoucou with the
Mexican deer goes back at least to Goldfuss 1817: 1122 (in von Schreber and
Goldfuss ca 1799-1817). The type antler rack of Ceruus mexicanus had been
illustrated by Pennant and by Hamilton Smith as the Mexican deer and could
readily have been seen by Brookes since it was in the collection of the British
Museum, London (Osgood 1920). Gray (1852: 228, 230) listed Elephalcedae [sic] in
the synonymy of Cariacus (= Odocoileus) and Elephalces [sic] mexicanus in the
synonymy of Ceruus mexicanus Gmelin. Thomas (1895: 193) listed Elephalces [sic]
as one of the many generic names given to American deer. Otherwise Elaphalces
mexicanus and the Elaphalcedae have been ignored in compendia. Elaphalces is a
senior synonym of Odocoileus, not oí Blastocerus as tentatively suggested by Grubb
(1993), but through lack of use is not valid. Elaphalcedae is a synonym of
Odocoileinae but can not have priority over the latter in view of Article 40 (b)
(ICZN 1985).
Subulidae Brookes, 1828: 62, is based on Subulus Brookes, loc. cit., as a
subfamily of Cervidae, so its suffix is not now regarded as properly formed. The
genus included S. Americanus, the 'Brocket' of 'New Jersey' = Ceruus americanus
Erxleben, 1777 = Odocoileus uirginianus, and S. spinosus = Ceruus spinosus Gay
and Gervais, 1846 = O. u. cariacou (Boddaert, 1784) but no type has been
Nomenclature of living and fossil deer
293
designated. Subulus Brookes is clearly a synonym of Odocoileus and though senior
to the latter, is not valid through lack of use. Subulus may seem to be an
emendation of Subulo Hamilton Smith, 1827, but the latter is a synonym of
Mazama Rafinesque, 1817, with type Cervus rufus Illiger, 1811 = Mazarna
americana (Erxleben, 1777).
Mazamadae Brookes, 1828: 62, was erected as a subfamily of Cervidae, with type
genus Mazama, by definition, and with improperly formed suffix. Brookes' Mazama
included only the Virginian 'Mazamme' Mazama Virginiana = Odocoileus virginianus (Zimmermann, 1718). Mazama in this context is therefore Mazama Hamilton
Smith, 1827, not Mazama Rafinesque, 1817. Hamilton Smith assigned what are
now Odocoileus species to Mazama, and true Mazama species to Subulo, both
treated as subgenera oí Cervus. Mazamadae Brookes is a synonym of Odocoileinae
but does not have priority over the latter in view of Article 40 (b) (ICZN 1985).
Mazaminae Kraglievitch, 1932: 426, with type genus Mazama Rafinesque by
definition and including also Pudu, is a junior homonym of Mazamadae Brookes
and hence is preoccupied.
Neocervinae Carette, 1922: 442, was coined to include Rangifer and genera of
deer confined to the Americas and was used subsequently by Flerov (1952) and
Banfield (1961), among others. It is not an available name as it is not based on a
genus-group name [Hershkovitz 1982: 5, ICZN 1985: Article 11 (f) (i) (1)], yet
continues to be employed in a formal sense (Vislobokova 1980, Kalandadze and
Rautian 1992: 127; the latter are cited by McKenna and Bell 1997: 429 as the
authors of Neocervinae but this is not the case).
Odocoileinae Pocock, 1923: 204, was proposed to include all genera of deer
confined to the Americas with the exception of Pudu. Elaphalcedae, Mazamadae
and Subulidae, all of Brookes, 1828, might appear to have priority over Odocoileinae
but have never been used by other authors. A family-group name with stem
Odocoile- is the senior available name for the endemic American genera of deer.
Simpson (1945) returned to Pocock's (1911) original concept of a subfamily
including all telemetacarpal Cervidae, but he called it Odocoileinae rather than
Capreolinae. Other authors followed this lead, though omitted the antler-less
Hydropotes (Ellerman and Morrison-Scott 1951, Anderson and Knox-Jones 1984,
Groves and Grubb 1987, Grubb 1993). But in this broader context, 'Odocoileinae' is
preoccupied by Capreolinae, Alceinae and Rangiferinae, all of Brookes, 1828.
Capreolinae should take priority according to Article 23 (d) (ICZN 1985) and
Pocock (1911: 971). Family-group names based on Capreolus, Alces and Rangifer
are all in use. As a name for all antlered telemetacarpal deer, Odocoileinae is a
junior synonym and cannot be employed in this sense, short of a ruling by the
Commission. Viret (1961) cited 'Odocoileini Simpson' but Simpson (1945) authored
the change in rank, not the name.
Cervus americanus is a name that has been coined on more than one occasion
and is mentioned here because references to the white-tailed or Virginian deer,
Odocoileus virginianus, in the older literature as O. americanus or Mazama
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P. Grubb
americana (the latter now the accepted name of a completely different species),
may not otherwise be understood. There are four relevant names:
(1) Cervus americanus Erxleben, 1777: 512, is now generally regarded as an
unavailable senior synonym of Odocoileus virginianus.
Under the heading of
Cervus dama, Erxleben (loc. cit.) wrote 'Differente vere americanus vti [uti]
Pennanto videtur?' followed by a description and references to early accounts of the
Virginian deer with a distribution including Virginia and Carolina. The name has
been written 'Cervus dama americanus', but should probably be cited as 'Cervus
americanus'. Allen (1900) thought americanus was an ordinary word, not a name,
but it was italicised. Osgood (1903) did not accept it as a name. Thomas (1913)
thought it was not a name for the very reason that it was italicised: Erxleben did
not italicise his scientific names (but Moschus americanus on page 317 is one
exception). Authors who considered brockets and Virginia deer congeneric have
called the latter Mazama americana (see Lydekker 1898).
(2) Moschus americanus Erxleben, 1777: 324, is now Mazama americana, the
accepted name for the red brocket and is likely to be confused with Cervus
americanus Erxleben in listings of synonyms and citations.
(3) Cervus americanus Clinton, 1822, is now Alces alces americanus.
(4) Cervus americanus Harlan, 1825, understood to be preoccupied, is now
Cervalces scotti (Lydekker, 1898).
From the above, it should be apparent that the name Mazama
americana
(Erxleben, 1777) as used in the literature has two separate origins and has been
applied to two quite separate species. Though Erxleben's name Cervus americanus
has long been abandoned, no ruling on its status has been made by the Commission
and technically it remains a senior homonym of Cervus americanus Clinton. It
might seem to threaten the stability of Moschus americanus Erxleben if Odocoileus
were once again regarded as a junior synonym of Mazama. This is not a wholly
hypothetical issue: authors who have considered that brockets and the Virginia
deer are congeneric published their views not only in the last century (Lydekker
1898), but also more recently (Haltenorth 1963). It is therefore desirable to ensure
that a ruling is made on the availability of Cervus americanus Erxleben.
Hippocamelus Leuckart, 1816: 23, based on Hippocamelus dubius Leuckart =
Equus bisulcus Molina, 1782, the huemal of Chile, would be the senior synonym if
all strictly American genera of deer were combined in one genus. With the
exception of Pudu, these genera were synonymized in Mazama by Lydekker (1898)
who used Xenelaphus Gray, 1869, instead oí Hippocamelus as the subgenus for the
huemals. Once he had appreciated that Hippocamelus must have ultimate priority,
he split the group into four genera to avoid 'the use of that highly objectionable
term' (Lydekker 1915: 155). This is not a wholly forgotten issue. More recently,
Haltenorth (1963) treated Blastóceros (= Ozotoceros), Hippocamelus, Mazama and
Pudu as subgenera of Odocoileus (including Blastocerus), ignoring priority once
again.
Nomenclature of living and fossil deer
295
Mazama Rafinesque, 1817, was named earlier than Odocoileus Rafinesque,
1832. It becomes a senior synonym of the latter when the two are combined, as was
done by Haltenorth (1963), who however ignored the priority. Mazama gouazoubira,
as usually quoted, is an unjustified emendation. The original citation is Cervus
gouazoupira G. Fischer, 1814: 465. Though Cabrera (1961: 338) regarded this as a
lapsus, there is no indication that the original spelling was other than what the
author intended (Grubb 1993: 391). The vernacular is 'gouazoú-birá' and A. L.
Gardner has petitioned the ICZN to validate the emendation (Medellín et al. 1998).
Mazama Hamilton Smith, 1827: 314, is preoccupied by Mazama Rafinesque,
1817. No type has been designated but it includes taxa now assigned to Blastocerus
dichotomus (Illiger, 1811), the marsh deer; Ozotoceros bezoarticus (Linnaeus,
1758), the pampas deer; Odocoileus virginianus and O. hemionus (Rafinesque,
1817), the mule deer.
Dorcelaphus Gloger, 1841: 140, with Cervus virginianus Zimmermann, 1780,
type by subsequent designation, is hence a junior subjective synonym of Odocoileus
Rafinesque, 1832. Dorcelaphus was coined to include Cervus campestris = Ozotoceros
bezoarticus, C. paludosus Desmarest, 1822 = Blastocerus dichotomus, C. ( =
Odocoileus) virginianus, C. macrourus = O. v. macrourus (Rafinesque, 1817) and
C. macrotis Say, 1823 = O. hemionus. Thomas (1895: 193) selected the type of
Dorcelaphus as Cervus virginianus, acting as first reviser. Subsequently, to replace
'Blastóceros [sic] Gray, 1850', Knottnerus-Meyer (1907) used Dorcelaphus as a
subgenus of Elaphus, including only D. dichotoma Illiger. He assigned the other
species included in Dorcelaphus by Gloger to other genera. Were it not for the
action of Thomas (1895), Dorcelaphus would be available as the generic name for
the marsh deer (Blastocerus dichotomous) since it predates Blastocerus Wagner.
Simpson (1945: 154) mistakenly thought that Dorcelaphus might have to replace
Blastocerus and more recently the marsh deer was cited as 'Dorcelaphus dichotomus' by Bubenik (1990: 70).
Blastocerus Gray, 1850a: 68 and Blastoce?'os Fitzinger, 1860: 176 are the correct
generic names of the marsh deer and pampas deer according to Hershkovitz's
(1958) review. It is undesirable to have virtual homonyms, which have been
thoroughly confused, applying to related but distinct deer, and therefore there is a
need to reconsider the situation. What Hershkovitz (1958) regarded as unitalicised
'Blastocerus' was, he says, coined by Wagner (1844: 366) as 'a diagnostic term for a
species group contained within the subgenus Elaphus' of genus Cervus and hence
was unavailable. Included in the group were Cervus paludosus = Blastocerus
dichotomus, C. campestris = Ozotoceros bezoarticus, and C. macrotis = Odocoileus
hemionus. Subsequent to Wagner's work, Gray (1850a: 68) used the name
Blastocerus including only Cervus paludosus and Fitzinger (1860: 176) in t u r n
employed Blastóceros for Cervus campestris (= Ozotoceros bezoarticus). The
species-group names paludosus and campestris were regarded as types of genera by
monotypy in each case and each generic name was regarded by Hershkovitz (1958:
15) as the first available name, other than 'Cervus', for each species. Each author
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could be regarded as making available the apparently unavailable 'Blastocerus' of
Wagner. Gray and Fitzinger acted independently and used different spellings and,
by monotypy, different types of genera, though neither stated that he had made an
unavailable name available or designated a type species. As long as Wagner's name
is unavailable, Blastocerus Gray and Blastóceros Fitzinger are to be treated as quite
independent entities in spite of their virtual homonymy. Blastóceros could not be
regarded as an unjustified emendation of Blastocerus. While indeed there is no
indication in the text of his work that Fitzinger adopted the name Blastóceros from
either Wagner (1844) or Gray (1850a) (Hershkovitz 1958: 14), both Fitzinger and
Gray were aware of Wagner's account, as is perfectly clear from their other
publications, where Wagner is cited. Gray (1850b, 1852, 1872) listed Blastóceras
paludosus and B. campestris, while in turn Fitzinger (1873, 1879) mentioned
Blastóceros paludosus and a trio of species (Blastóceros campestris, B. azarae, B.
comosa) all now regarded as representing the pampas deer. To have Blastocerus
Gray and Blastóceros Fitzinger as independent genera does not contribute to
stability, especially as citations of each name have often been mis-spelt as the other
(Ameghino 1891, Knottnerus-Meyer 1907, Lydekker 1915, Cabrera 1961) and
particularly because it is patent that Gray and Fitzinger each used a single genus to
encompass both the marsh deer and the pampas deer.
We can be rescued from a situation which must seem unfortunate to many
zoologists. It seems that Hershkovitz (1958) can now be regarded as mistaken in his
interpretation of Wagner (1844). According to Article 10 (e) (ICZN 1985) a 'secondary
(or further) subdivision [of a genus], is deemed to be a subgeneric name even if the
division is denoted by a term such as "section" or "division"'. Therefore, Blastocerus
Wagner, with type by subsequent designation (this paper) Ceruus paludosus ( =
Blastocerus dichotomus), can stand as a genus-group name. A formal statement
that Cervus paludosus should be selected as the type of Blastocerus Wagner cannot
be traced, so this designation is made herewith else technically Blastocerus Wagner
could become a junior synonym of Odocoileus or a senior synonym of Ozotoceros,
outcomes implicitly posed by Simpson (1945) and which are to be avoided.
Since Blastocerus Wagner is available, Fitzinger's use of Blastóceros does after
all amount to an unjustified emendation, an alteration of the Latinised version of
the Greek (Blastocerus) to a transliterated Greek (Blastóceros) which 'is available
with its own author and date and is a junior or objective synonym of the name in its
original spelling' [ICZN 1985: Article 33 (b) (iii)]. As the author 'replaces a
previously established genus-group name by an unjustified emendation ... both the
prior nominal taxon and its replacement must have the same type species' [ICZN
1985: Article 66 (h)]. Blastocerus Wagner and Blastóceros Fitzinger are objective
synonyms and the latter is no longer available as a senior synonym of Ozotoceros.
Eucervus Gray, 1866: 338, type Cervus macrotis = Odocoileus hemionus by
subsequent designation of Miller (1924: 484), and also including C. columbianus
Richardson, 1829 (= 0. h. columbianus), is a senior homonym of Eucervus Aclogue,
1899, and cannot be a type genus of Eucervidae Bubenik, 1990.
Nomenclature of living and fossil deer
297
Ozotoceros Ameghino, 1891: 243, with type by original designation 'Blastoceros
[sic] campestris Gray = Cervus campestris Cuvier' (correctly, Ceruus campestris in
the sense of Wied 1826, and other authors, not of Cuvier, = C. bezoarticus
Linnaeus, 1758). It is the earliest generic name based on the pampas deer (pace
Hershkovitz 1958: 14) since as explained above, Blastoceros Fitzinger is not
available for this species. Cervus campestris F. Cuvier, 1817, was long regarded as a
name for the pampas deer, but Cuvier's name is a synonym of Odocoileus
virginianus cariacou (Boddaert, 1784) according to Cabrera (1943: 21, 1961: 325).
Cabrera (1961: 330) implies that the first use of Cervus campestris in place of
Cervus bezoarticus Linnaeus was by Wied in 1826 [not seen].
Pudu puda (Capra puda Molina, 1782) has been universally cited as lPudu
pudu', an unjustified emendation (Hershkovitz 1982: 64-65).
Procervus de Blainville, 1840: 392, a renaming of Procerus de Serres, is a senior
homonym of Procervus Hodgson, 1847 [= Cervus (Rucervus) Hodgson, 1838].
Procerus de Serres, 1832-1834 (according to Sherborn 1929: 5159) or 1838
(according to Palmer 1904: 566 and McKenna and Bell 1997), with type by
monotypy P. cariboeus de Serres = Rangifer tarandus (Linnaeus, 1758), is in t u r n
preoccupied by Procerus Megerle, 1821 (Coleoptera; Sherborn 1929) and both are
said to be preoccupied by a still earlier name, Proceros Rafinesque, 1820, a genus of
fish (Palmer 1904, Sherborn 1929), though this would no longer be correct under
the Code.
P l e s i o m e t a c a r p a l deer, i n c l u d i n g Muntiacus
and allies
Plesiometacarpi Brooke, 1878: 897, is not an available name, as indicated under
Telemetacarpi. It was recognised as equivalent to Cervinae by Pocock (1911: 971),
which in any case would have priority.
Stylocerinidae Brookes, 1828: 62, was erected for Diopplon Brookes, loc. cit., as a
subfamily of Cervidae, but presumably is based on Stylocerus Hamilton Smith,
1827, and if so the prefix as well as the suffix is improperly constructed. Stylocerinidae is not available because it does not satisfy Articles 11 (f) (i) (1) or 64 (ICZN
1985): a family-group name must have a type genus that the author considers to be
valid. By citing Diopplon, Brookes failed to indicate the validity of Stylocerus.
Cervulinae Sclater, 1870: 115, is a senior synonym of Muntiacinae Knottnerus-Meyer, 1907, but as the latter has been used generally, Cervulinae cedes this
seniority for the purposes of synonymy [Article 40 (b) of the Code] and in any case
has been placed on the Official Index of Rejected and Invalid Family-Group Names
in Zoology (ICZN 1959), though it has continued to be used (Gromova 1962),
together with tribal and subtribal names Cervulini and Cervulina (Kalandadze and
Rautian 1992: 126).
Elaphodinae Knottnerus-Meyer, 1907: 15, for Elaphodus Milne-Edwards, 1871,
is here declared to be junior to Muntiacinae when the two are combined.
298
P. Grubb
Muntiacinae Pocock, 1923: 207 has been placed on the Official List of Family-Group Names in Zoology (ICZN 1959) but is pre-dated by Muntiacinae Knottnerus-Meyer, 1907: 14, 97, also based on Muntiacus Rafinesque, 1815.
Diopplon Brookes, 1828: 62, was erected as a new genus for Ceruus muntjak
Zimmermann, 1780. It has been omitted from checklists (Ellerman and Morrison-Scott 1951, Corbet and Hill, 1992, Grubb 1993) and is preoccupied by Muntiacus
Rafinesque, 1815, Ceruulus Blainville, 1816, and Stylocerus Hamilton Smith, 1827.
Muntiacus Rafinesque, 1815, has been validated by Opinion 460 (ICZN 1957)
with genotype Cervus muntjak Zimmerman, 1780. The change from Muntiacus feae
(Thomas and Doria, 1889) to M. feai was an emendation made on the advice of
Professor Tortonese (Grubb 1977), but as noted in Grubb (1993: 389) it was
unjustified [ICZN 1985: Article 31 (a)], the names of the Italians Leonardo Fea and
Giacomo Doria having been consistently treated as if first declension Latin with
genitives 'Feae' and 'Doriae' (eleven species-names cited by Wilson and Reeder
1993) in accord with Article 31 (a).
P l e s i o m e t a c a r p a l deer: Cervus
and allies
Axidae Brookes, 1828: 61, was erected as a subfamily of Cervidae based on Axis
Hamilton Smith, 1827, so would now be regarded as having been formed with the
incorrect suffix.
Platycerinidae Brookes, 1828: 61, was erected as a subfamily of Cervidae, presumably based on Platyceros Zimmermann, 1780, a junior synonym of Dama Frisch,
1775, to include only Dama. Platycerinidae is improperly formed both in suffix and
prefix, and is not available for the reasons cited above under Stylocerinidae.
Rusadae Brookes, 1828: 62, was erected as a subfamily of Cervidae, based on
Rusas, presumably an incorrect spelling of Rusa Hamilton Smith, 1827. It is an
available name but with incorrectly formed suffix and prefix. Gray (1852: 201)
corrected it to Rusinae, to include Axidae and Stylocerinidae Brookes and the
genera Panolia, Rucervus, Rusa, Axis, Hyelaphus and Cervulus. Blyth (1863: 149)
also used the correctly formed Rusinae as a subfamily of Cervidae to include the
same genera as Gray with the exception of Cervulus.
Elaphidae Brookes, 1828: 61, was erected as a subfamily of Cervidae, based on
Elaphus Hamilton Smith, 1827, so would now be regarded as improperly formed. It
was corrected to Elaphinae by Gray (1852: 193). Elaphus is an objective synonym of
Cervus, so Elaphidae is a junior objective synonym of Cervidae.
Megaloceridae Brookes, 1828: 61, was coined as a subfamily of Cervidae with
type genus Megalocerus by definition. It not only has an incorrectly formed suffix
but as Megalocerus is to be regarded as an emendation of Megaloceros (ICZN 1989:
Opinion 1566), it is now seen to have an incorrect stem of the generic name and
should be spelt 'Megalocerotinae' by analogy with Rhinoceros and Rhinocerotidae
or Strepiceros and Strepsicerotinae: it is a transliteration of the Greek, not a
Nomenclature of living and fossil deer
299
Latinised version of the Greek, so should adopt a Greek stem [ICZN 1985: Article
29 (b), Appendix B, Table II]. Megacerini Viret, 1961: 1018, based on Megaceros
Owen, 1844, is a junior synonym of Megalocerotinae Brookes, 1828.
Eucervidae Bubenik, 1990: 109, is not an available name [ICZN 1985: Articles 11
(f) (i), 132 (a) (i)]. No type genus is cited and while the family could presumably be
based on Cervus (Eucervus) Aclogue, no reference to this nominal subgenus is
made. In any case, Eucervidae Bubenik is preoccupied by Cervidae Goldfuss, 1820.
Cervus (Eucervus) Aclogue, 1899: 71 (not of Gray, 1866), with type Cervus elaphus
Linnaeus, 1758, by monotypy, is an objective synonym of Cervus Linnaeus.
Axis javanicus (Cervus axis javanicus von Koenigswald, 1933: 65), is preoccupied
javanicus)
by Cervus javanicus 'Gmelin, 1766' (that is, Osbeck, 1765 = Tragulus
according to Kretzoi (1947: 286) and was renamed as Axis sunda Kretzoi, 1947.
Axis japonicus [Cervus (Axis) japonicus Otsuka, 1967: 279] is preoccupied by
Cervus (Hippelaphus) japonicus Sundevall, 1846: 178 (= Cervus nippon Temminck,
1838). It is here renamed Axis kyushuensis nomen novum.
Cervus vulgaris montanus Botezat, 1903: 155, is treated as a subspecies (the
Carpathian red deer, C. elaphus montanus) with synonym C. v. campestris Botezat,
1903: 154, by Heptner et al. (1961) but both names are doubly invalid, being
nomina nuda and preoccupied by Cervus macrotis var. Montanus Caton, 1881 ( =
Odocoileus hemionus) and Cervus campestris F. Cuvier, 1817 (= Odocoileus virginianus cariacou), respectively. Cervus vulgaris is an objective junior synonym of
C. elaphus Linnaeus, 1758. Cervus elaphus sibiricus Severtzov, 1872, has been
thought to be preocupied by C. sibiricus Schreber, 1784 (a synonym of Rangifer
tarandus), but the latter is not regarded as a binomial term and so cannot displace
Severtzov's name (Ellerman and Morrison-Scott 1951: 376, Heptner et al. 1961:
160).
Cervus (Deperetia) Shikama, 1936b: 251, for Cervus (confer Anoglochis) praenipponicus Shikama, 1936a: 170, is preoccupied by Deperetia Teppner, 1921 (Mollusca),
and Deperetia Schaub, 1923 (Mammalia, Bovidae) and has been replaced by Nipponicervus Kretzoi, 1941: 350, with the same type, which is in t u r n a junior synonym of
Bohlinella Palmer, 1939: 110, coined independently to replace Deperetia Shikama.
Yet Deperetia Shikama has continued in use (Otsuka 1967, Vislobokova 1990).
Elaphus Hamilton Smith, 1827: 307, is an objective synonym of Cervus, both
having Cervus elaphus as type, but it was employed by Brookes (1828) instead of
Cervus and is still being used in a subgeneric sense in place of Cervus (Cervus)
(Dong, 1993).
Procervus Hodgson, 1847: 689, with type by monotypy Cervus
dimorphe
Hodgson, 1843: 897 = Cervus (or Rucervus) duvauceli Cuvier, 1823, has been noted
by Palmer (1904: 567) and in palaeontological papers (Dietrich 1938, Czyżewska
1968) but has been omitted from checklists (Lydekker 1915, Ellerman and Morrison-Scott 1951, Corbet and Hill 1992, Grubb 1993). It is preoccupied by Procervus de
Blainville, 1840, and is a senior homonym of Procervus Alexejev, 1913, and a junior
synonym of Rucervus Hodgson, 1838.
300
P. Grubb
Pseudodama Azzaroli, 1992: 4, with type Cervus nestii Azzaroli, 1947, by
original designation, was coined to include the other Villafranchian deer C.
pardinensis Croizet and Jobert, 1828, C. rhenanus Dubois, 1904, C. perolensis, P.
lyra Azzaroli, 1992, and P. farnetensis Azzaroli, 1992. It is therefore preoccupied by
Cervus (Metacervocerus) Dietrich and perhaps also by Cervus (Praeelaphus) Portis.
Cervus (Metacervocerus) Dietrich, 1938: 265, has Cervus pardinensis as type by
original designation with presumed synonyms C. etueriarum, C. perrieri, C.
issiodorensis (all of Croizet and Jobert) and C. rhenanus. Cervus (Praeelaphus)
Portis, 1920: 133 was coined to include C. arvernensis Croiset and Jobert, 1828, C.
etueriarum and C. perrieri (with synonyms issiodorensis and pardinensis). It may
therefore be a senior synonym of Cervus (Metacervocerus) Dietrich, and of Pseudodama Azzaroli, but no type has been designated and the name has achieved no
currency.
Alee Blumenbach, 1799: 697, with type A. gigantea Blumenbach, loc. cit. ( =
Megaloceros giganteus) would be preoccupied by Alee Frisch, 1775, for Cervus alces
(ICZN 1926: Opinion 91), but Frisch, 1775 is unavailable (ICZN 1950): Dama
Frisch is an exception, having been recognised as an available name (ICZN 1960).
So Alee Blumenbach is available but in view of its lack of use and its similarity to
Alces, associated also with a deer having large palmate antlers, it should be
suppressed. Megaceros Owen, 1844, is a junior synonym of Megaloceros Brookes,
1828, and can not be used as a subgenus (Geist 1999: 122) when the types of both
genera are believed to represent the same species.
Discussion
This paper has reviewed problems concerning cervid nomenclature including
previously evaluated but neglected instances of availability, priority, synonymy,
homonymy, emendation and authorship. Unresolved or contentious issues have
been addressed and family-group names in Cervidae have been reviewed. Pliocervinae Khomenko, Telemetacarpi, Plesiometacarpi, Neocervinae Carette and
Eucervidae are unavailable. Family-group prefixes and suffixes are to be corrected
if inappropriately formed, and changes in rank can occur, following the Principle of
Co-ordination (ICZN 1985: Article 36). Family-group names should retain their
original authors and are not attributable to those who altered their rank or made
justifiable emendations of original spellings: names based on Capreolus, Alces and
Rangifer are hence attributable to Brookes, 1828, not to Pocock, Simpson or others.
The widely used emendation Alcinae is due to Blyth, not Jerdon, but has been
emended definitively to Alceinae (ICZN 1977b: Opinion 1081). Capreolinae, Alceinae
and Rangiferinae take precedence over Odocoileinae when the taxa are combined,
contrary to common practice. Alceinae has priority over Rangiferinae where relevant
(here designated). Brookes (1828) - though an available work (ICZN 1977a) includes overlooked and incorrectly formulated family-group names Stylocerinidae
(senior to Muntiacinae; unavailable); Platycerinidae (for Dama; unavailable);
Nomenclature of living and fossil deer
301
Mazamadae (unavailable senior synonym of Odocoileini); Elaphalcedae and
Subulidae (senior synonyms of Odocoileinae but have ceded their seniority); Axidae
(available); Rusadae (has been emended to Rusinae; available); and Elaphidae
(objective synonym of Cervidae). Mazaminae Kraglievitch is preoccupied by
Mazamadae Brookes: they are not synonyms. Cervulinae had been made unavailable,
but is still being used. The author of Muntiacinae is Knottnerus-Meyer, 1907, not
Pocock, 1923, and this taxon includes Elaphodinae as a junior synonym (here
designated). Megacerinae Viret is preoccupied by Megalocerinae Brookes, which in
t u r n is here emended to Megalocerotinae. Holometacarpal deer have been allocated
to Pliocervinae Symeonidis, 1974, including Cervavitus and Pliocervus (for synonyms see Table 1). Metadicrocerus (junior synonym and available replacement for
preoccupied Proceruus Alexejev), problematic senior homonym Cervus americanus
Erxleben (senior synonym of Odocoileus virginianus and senior homonym of Alces
alces americanus), Elaphalces and Subulus (senior synonyms of Odocoileus),
Diopplon (junior synonym of Muntiacus), Proceruus Hodgson (junior synonym of
Ruceruus), and Alee Blumenbach (senior synonym of Megaloceros) have evaded the
attention of compilers and the senior synonyms will have to be suppressed.
Homonyms of nomenclatural significance include Mazama Rafinesque, 1817, and
Mazama Hamilton Smith, 1827; Euceruus Gray, 1866, and Euceruus Aclogue, 1899;
Proceruus de Blainville, 1840, Proceruus Hodgson, 1847, and Proceruus Alexejev,
1913; Ceruus americanus Erxleben, 1777, C. americanus Clinton, 1822, and C.
americanus
Harlan, 1825; C. jauanicus Osbeck, 1765, and C. jauanicus von
Koenigswald, 1933; C. japonicus Sundevall, 1846, and C. japonicus Otsuka, 1967;
C. montanus Caton, 1881, and C. montanus Botezat, 1903; C. campestris F. Cuvier,
1817, and C. campestris Botezat, 1903. Near homonyms include Subulo Hamilton
Smith, 1827, and Subulus Brookes, 1828; and Alee Blumenbach, 1799, and Alces
Gray, 1821. Near homonyms Blastocerus Wagner, 1844, and Blastóceros Fitzinger,
1860, are here shown to be synonyms: Ceruus paludosus Desmarest is confirmed as
the lectotype of Blastocerus Wagner, which is an available name of which Blastóceros Fitzinger is an unjustified emendation, not a senior synonym of Ozotoceros.
Cervus americanus Erxleben (white-tailed deer) and Moschus americanus Erxleben
(red brocket) have both been cited in the older literature under the name Mazama
americana, leading to potential confusion in tracing synonymy. Dorcelaphus is a
junior synonym of Odocoileus, not a senior synonym of Blastocerus. Elaphalces is
not a synonym of Blastocerus either. Like Pudu pudu, Mazama
gouazoubira
remains an unjustified emendation (correctly, M. gouazoupira). Order of seniority
among odocoileine genera (Hippocamelus > Mazama > Odocoileus > Blastocerus
> Pudu > Ozotoceros) should be recognised if any are to be synonymised.
Muntiacus feae should not be emended to M. feai. Axis kyushuensis nomen novum
is proposed for Ceruus japonicus Otsuka, preoccupied. Deperetia Shikama, though
known to be preoccupied, is still being used for Cervus (Bohlinella)
praenipponicus.
Cervus (Elaphus), also in use, is invalid since it is an objective synonym of Cervus.
The name for the Carpathian red deer (Cervus elaphus montanus Botezat, 1903) is
302
P. Grubb
Table 1. Interim classification of deer, Cervidae, citing all extant genera, extinct genera mentioned in
the text, and other well-known extinct genera. Authors and dates of names are given only for
family-group names and for homonymous genera. Unavailable family-group names are not included.
The classification is modified from Groves and Grubb (1987) and McKenna and Bell (1997). The latter
should be consulted for a more complete list of extinct genera and synonyms.
Family Cervidae Goldfuss, 1820
Subfamily Hydropotinae Troussart, 1898
Hydropotes
Subfamily Lagomerycinae Pilgrim, 1941
Tribe Lagomerycini Pilgrim, 1941
tLagomeryx,
tProcervulus
Tribe Dicrocerini Simpson, 1945
tDicrocerus, tEuprox, tHeteroprox, etc.
Subfamily Pliocervinae Symeonidis, 1974
tCervavitus (synonyms: tCervocerus, tDamacerus, tMetadicrocerus,
tProcervus Alexejev, 1913, not of de Blainville, 1840), tPliocervus
(synonyms: tCtenocerus Kretzoi, 1941, not of Dahlbom, 1846;
t Ctenocervus)
Subfamily Capreolinae Brookes, 1828
Tribe Capreolini Brookes, 1828
tProcapreolus,
Capreolus
Tribe Alceini Brookes, 1828
t Cerualces, Alces
Tribe Odocoileini Pocock, 1923 (synonyms: Elaphalcedae, Subulidae and
Mazamadae, all of Brookes, 1828; Mazaminae Kraglievitch, 1932)
tPavlodaria, tBretzia, Mazama Rafinesque, 1817 (synonym: Subulo),
Pudu, Hippocamelus, tAgalmaceros, tNauahoceros, Ozotoceros,
Blastocerus, Odocoileus (synonyms: Mazama Hamilton Smith, 1827,
not of Rafinesque, 1817, Elaphalces, Subulus, Dorcelaphus,
Eucervus
Gray, 1866), tMorenelaphus, etc.
Tribe Rangiferini Brookes, 1828
Rangifer (synonyms: tProcerus, tProcervus de Blainville, 1840)
Subfamily Cervinae Goldfuss, 1820
Tribe Muntiacini Knottnerus-Meyer, 1907 (synonyms: Cervulinae Sclater,
1870; Elaphodinae Knottnerus-Meyer, 1907)
Elaphodus, Megamuntiacus, tEostyloceros, Muntiacus (synonyms:
Cervulus, Stylocerus, Diopplon), etc.
Tribe Cervini Goldfuss, 1820 (synonyms: Axidae, Rusadae, Elaphidae,
Megaloceridae, all of Brookes, 1828; Megacerini Viret, 1961)
tCroizetoceros, Axis, Hyelaphus, Wohlinella (synonyms: 1 Deperetia
Shikama, 1936, not of Teppner, 1921, or Schaub, 1923,
tNipponicervus), tMetacervocerus (synonym: tPseudodama) possibly
preoccupied by tPraeelaphus, Rusa, Rucervus (synonym: Procervus
Hodgson, 1847, not of de Blainville, 1840), Cervus (synonyms: Sika,
Elaphus, Eucervus Aclogue, 1899, not of Gray, 1866), Elaphurus,
tEucladoceros, Przewalskium, tArvernoceros, Dama (synonym:
Platyceros), tPraemegaceros, tMegaceroides,
tMegaloceros
(synonyms: fAlce Blumenbach, 1799, not of Frisch, 1775, but
available; tMegaceros), tSinomegaceros, etc.
Nomenclature of living and fossil deer
303
a preoccupied nomen nudum. Pseudodama is preoccupied by Metacervocerus and
possibly Praeelaphus. This paper does not have space to attempt a review of the
evidence for a revised classification of the Cervidae, but a classification is provided
in Table 1, which summarises the author's provisional views and includes what is
believed to be valid nomenclature for genera, tribes and subfamilies. Other workers
may of course prefer to change the rank of some taxa or to place some genera in
different subfamilies or tribes.
Acknowledgements: I am grateful to Adrian Lister and two anonymous referees for their constructive
comments on earlier versions of this paper.
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Received 3 December 1999, accepted 21 February
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